1. Max Planck Institute for Limnology, August Thienemann Strasse 2, 24306 Ploen, Germany
2. National Environment Research Council Centre for Population Biology, Imperial College at Silwood Park, Ascot, Berkshire SL5 7PY, England, United Kingdom
3. Department of Biological Sciences and National Environment Research Council Centre for Population Biology, Imperial College at Silwood Park, Ascot, Berkshire SL5 7PY, England, United Kingdom